A peer-reviewed open-access journal PhytoKeys 29: 17–62Diversity (2013) and levels of endemism of the Bromeliaceae of Costa Rica... 17 doi: 10.3897/phytokeys.29.4937 CHECKLIST www.phytokeys.com Launched to accelerate biodiversity research Diversity and levels of endemism of the Bromeliaceae of Costa Rica – an updated checklist Daniel A. Cáceres González1,2, Katharina Schulte1,3,4, Marco Schmidt1,2,3, Georg Zizka1,2,3 1 Abteilung Botanik und molekulare Evolutionsforschung, Senckenberg Forschungsinstitut Frankfurt/Main, Germany 2 Institut Ökologie, Evolution & Diversität, Goethe-Universität Frankfurt/Main, Germany 3 Biodive rsität und Klima Forschungszentrum (BiK-F), Frankfurt/Main, Germany 4 Australian Tropical Herbarium & Center for Tropical Biodiversity and Climate Change, James Cook University, Cairns, Australia Corresponding author: Daniel A. Cáceres González ([email protected]) Academic editor: L. Versieux | Received 1 March 2013 | Accepted 28 October 2013 | Published 11 November 2013 Citation: González DAC, Schulte K, Schmidt M, Zizka G (2013) Diversity and levels of endemism of the Bromeliaceae of Costa Rica – an updated checklist. PhytoKeys 29: 17–61. doi: 10.3897/phytokeys.29.4937 This paper is dedicated to the late Harry Luther, a world leader in bromeliad research. Abstract An updated inventory of the Bromeliaceae for Costa Rica is presented including citations of representa- tive specimens for each species. The family comprises 18 genera and 198 species in Costa Rica, 32 spe- cies being endemic to the country. Additional 36 species are endemic to Costa Rica and Panama. Only 4 of the 8 bromeliad subfamilies occur in Costa Rica, with a strong predominance of Tillandsioideae (7 genera/150 spp.; 75.7% of all bromeliad species in Costa Rica). 124 species (62.6%) grow exclusively epiphytic, additional 59 spp. (29.8%) are facultative epiphytes. The most diverse genus is Werauhia, with 59 species (29.8% of the Costa Rican bromeliad flora), followed by Tillandsia with 40 species (20.2%) and Guzmania with 28 spp. (8.6%). Resumen Es presentada una lista de chequeo actualizada de 18 géneros y 198 especies de Bromeliaceae conocidas de Costa Rica. Por consiguiente, las bromelias endémicas de Costa Rica, ahora comprenden 32 especies. Adicionalmente, 36 especies tienen un endemismo compartido con Panamá. Sólo 4 de las 8 subfamilias actualmente reconocidas ocurren en Costa Rica con fuerte predominio de Tillandsioideae (7 géneros/150 spp.; 75.7% del total de especies de bromelias de Costa Rica). 124 especies (62.6%) crecen como epífita, 59 especies adicionales (29.8%) son epifitas facultativas. El género mejor representado es Werauhia, con 59 especies (29.8% de la flora de bromelias de Costa Rica), seguido por Tillandsia con 40 spp. (20.2%) y Guzmania con 28 especies (8.6%). Copyright Daniel A. Cáceres González et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 18 Daniel A. Cáceres González et al. / PhytoKeys 29: 17–61 (2013) Keywords Central America, epiphytism, life-form, systematic diversity Introduction Costa Rica is part of the hotspots of plant diversity in the Neotropics (Myers et al. 2000) with an estimated 8,249 vascular plant species (Hammel et al. 2004). Over the last century the number of scientifically documented plants for Costa Rica has increased considerably rendering the country one of the best studied in Mesoamerica. In Costa Rica, the Bromeliaceae belong to the families with highest species diversity in epiphytic habitats and thus contribute considerably to overall forest diversity. Bromeliaceae comprise 3,172 species and 58 genera (Luther 2008), which are grouped in the 8 subfamilies Brocchinioideae, Lindmanioideae, Tillandsioideae, Hechtioideae, Navioideae, Pitcarnioideae, Puyoideae and Bromelioideae (Givnish et al. 2011). All bromeliads are restricted to the Neotropics, with the one exception of Pitcairnia feliciana from West Africa. Based on the extensive revision of herbarium collections from Costa Rica, we pre- sent an updated checklist of Bromeliaceae for the country, and provide a brief analy- sis of systematic diversity, levels of endemism, distribution and life-form spectrum (epiphytic/terrestrial/saxicolous). Recent floristic work, in particular the revision of the diversity of Bromeliaceae of Panama (Cáceres González et al. 2011b) required the reas- sessment of endemicity of Costa Rican bromeliads. This led to considerable changes in the number of endemic bromeliad species recognized for Costa Rica. Materials and methods Herbarium collections of Bromeliaceae from 23 herbaria (B, C, CR, DUKE, F, FR, GH, INB, K, LI, MICH, MO, NY, PMA, RM, SCZ, SEL, TEX, US, USJ, UMO, WS and WU; abbreviations after Thiers 2008) were studied. Altogether 4,523 herbarium specimens of Bromeliaceae from Costa Rica were revised. For identification and extraction of biological information, the following referenc- es were used: Smith and Downs (1974, 1977, 1979), Utley (1994), Méndez-Estrada (1995), Grant (1995a, 1995b), Luther and Kress (1996), Grant and Morales (1996), Rossi et al. (1997), Morales (1999, 2000, 2003a, 2003b, 2003c, 2005, 2009), Morales and Alfaro (2003), Luther (2003), Cascante Marín et al. (2008), and Cáceres González et al. (2011a, 2011b). Further relevant references are given under the relevant species. All species are listed below with information on their distribution and life form. Species endemic to Costa Rica are marked with an asterisk *. The endemic species re- ported by Morales (2003c) are marked with a “1”. Species endemic to Costa Rica and Panama are marked with a “2”. Diversity and levels of endemism of the Bromeliaceae of Costa Rica... 19 Type specimens are annotated (holo = holotype, iso = isotype, lecto = lectotype, para = paratype). Additionally, a maximum of five revised herbarium collections are listed for each species, except in cases where the number of available collections was less than five. The presented checklist includes indigenous taxa and only one cultivated and nat- uralized taxon (Ananas comosus (L.) Merr.). Subspecies, varieties, and forms are not included in the list. Species erroneously reported for Costa Rica are listed separately. Synonyms (syn.) are only included if found in the recent literature. Nomenclature, generic delimitation and total species numbers for genera follow Luther (2008) and the International Plant Names Index (IPNI 2011). Results and discussion Bromeliaceae diversity in Costa Rica According to our studies, Costa Rica harbours 18 genera and 198 species of Bromeli- aceae, which represents 2.4% of the total angiosperm flora of the country (see species list [Appendix] and and Table 1). Thus, species diversity of the Costa Rican Bromeli- aceae is one of the richest among the Central American countries. Of the 3,172 species and 58 genera recognized for the whole family (Luther 2008), 6.2% of the species and 31% of the generic diversity are represented in Costa Rica. The increase in species reported for Costa Rica in the recent past (e.g. Utley (1994): 169 species; Luther (1995): 192 spp.; and Morales (2003c): 195 spp) can mostly be attributed to the discovery and description of new species (Luther and Kress 1996: Guz- mania herrerae H. Luther & W. J. Kress, G. scandens H. Luther & W. J. Kress; Grant and Morales 1996: Pitcairnia calcicola J. R. Grant & J. F. Morales; Morales 1999: Vriesea barii J.F. Morales, V. simulans J.F. Morales, V. haberii J.F. Morales, V. osaensis J.F. Mo- rales, V. tiquirensis J.F. Morales, V. vulcanicola J.F. Morales). Nevertheless, there were several discrepancies between previous studies of the bromeliad flora for Costa Rica (e.g. Luther 1995; Morales and Alfaro 2003; Morales 2003c), which we were able to resolve. For example, Luther (1995) added Tillandsia streptophylla Scheidw. ex C. Morren as new for Costa Rica. Our attempt to trace the specimens on which such report was based was not successful, and perhaps for this reason, Morales (2003c) refrained from including T. streptophylla in his list of bromeliads of Costa Rica. Nevertheless, we were able to verify the presence of the species based on recently collected specimens revised by us, thus T. streptophylla is again included in our checklist (see Appendix). According to Morales (2009), Aechmea penduliflora André, formerly reported for Costa Rica, does not exist in the country. Morales (2009) refers to misidentified col- lections (Espinoza 94, INB, MO; McPherson 8533, MO; Rueda & Mendoza 17152, MO), which represent A. angustifolia Poepp. & Endl. However, other specimens from Costa Rica revised by us represent A. penduliflora (see Appendix), thus we included the species in our list. 20 Daniel A. Cáceres González et al. / PhytoKeys 29: 17–61 (2013) Table 1. Bromeliaceae of Costa Rica: species richness and endemism. Number of % of bromeliad Number of endemic % of endemic Subfamily & genus species flora species species Tillandsioideae Catopsis 11 5.6 1 3.1 Guzmania 28 14.1 4 12.5 Mezobromelia 1 0.5 0 0.0 Racinaea 5 2.5 0 0.0 Tillandsia 40 20.2 2 6.3 Vriesea 6 3.0 2 6.3 Werauhia 59 29.8 18 56.3 Subtotal=7 150 75.7 27 84.5 Bromelioideae Aechmea 17 8.6 0 0 Ananas 2 1.0 0 0 Androlepis 1 0.5 0 0 Araeococcus 1 0.5 0 0 Billbergia 1 0.5 0 0 Bromelia 3 1.5 0 0 Greigia 2 1.0 0 0 Ronnbergia 1 0.5 0 0 Subtotal=8 28 14.1 0 0.0 Pitcairnioideae Pepinia 1 0.5 1 3.1 Pitcairnia 17 8.6 3 9.3 Subtotal=2 18 9.1 4 12.4 Puyoideae Puya 2 1.0 1 3.1 Subtotal=1 2 1.0 1 3.1 TOTAL=18 198 100.0 32 100.0 Morales (2003c) included Catopsis werckleana Mez in the synonymy of Catopsis nutans (Sw.) Griseb. However, we accept C.